li .2l

Reprinted from the Proceedings of the Pa. Academy of Science, Vol. 42, 1968

THE FAUNA OF BEAR MEADOWS, A BOREAL BOG IN CENTRAL PENNSYLVANIA

H. B. WHITE, III Graduate Department of Biochemistry Brandeis University Waltham, Massachusetts 02154

G. H. AND A. F. BEATTY P. 0. Box 281 State College, Pennsylvania 16801

ABSTRACT In this report are compiled all known Odon­ Fifty-eight of Odonata are re­ ata records for Bear Meadows through 1966. Since collecting has covered all months from corded as occurring at Bear Meadows, a May to November, the seasonal distribution of sphagnum bog in Central Pennsylvania many of the 58 species known from Bear Mea­ Seasonal distribution, relative abundance dows can be estimated. The data presented in and habitat preference are discussed. this paper are primarily the authors'; however, included are previously unpublished contribu­ tions of a number of collectors: J. Chemsak, INTRODUCTION C. Cook, T. W. Donnelly, S. W. Frost, J. Gil­ Bear Meadows is a 520-acre sphagnum bog lespie, J. G. Needham, R. A. Raff, J. M. Run­ nestled in the mountains of central Pennsyl­ ner and J. Schmidt. vania (Figs. 1, 2). It is 56 miles southwest of the nearest terminal moraine of Wisconsin LOCATION AND DESCRIPTION glaciation (1), the location of which makes it OF BEAR MEADOWS a very interesting place to study boreal organ­ Bear Meadows lies abov.t 7 miles south of isms at or near their southern limit of distribu­ State College in a protected mountain valley tion (2). Botanists are quite familiar with this 1824 ft above sea level. A semicircular moun­ bog because boreal species, such· as balsam fir tain ridge 500 ft above the valley surrounds and black spruce, grow here in association with the bog except for two openings to the north­ a flora more typical of Pennsylvania (3,4,5,6). east (Fig. 2). Access is by unpaved roads from Study of the of Bear Meadows has U.S. Routes 322 and 545. revealed a pattern of association similar to that Palynological studies indicate that Bear found with plants. Meadows was formed 10,000 years ago when In the summer of 1931, James G. Needham drainage from the area was blocked, creating a of Cornell University, one of the preeminent shallow lake (6,7). The normal progression of workers on dragonflies, was a visiting professor bog formation has proceeded to the point where at the Penn State Nature Camp, located at forest has invaded much of the bog's original what is now Alan Seeger State Forest Monu­ area leaving at present a one and one quarter ment in Huntingdon County. He went with mile open strip along peat-stained Sinking a party of students and faculty to visit Bear Creek. On either side of this meandering Meadows, then difficult of access, because it stream, thick mats of sphagnum and peat sup­ was said to be an interesting place although port marsh grasses, sedges, leatherleaf and ex­ little-studied biologically. He caught speci­ tensive stands of high bush blueberries. The mens of Leucorrhinia hudsonica and Somato­ invading forest consists primarily of black chlora elongata, observed the unco=on tree spruce, hemlock, balsam fir, red maple, black Prunus alleghaniensis, and came back to camp gum and a dense undergrowth of rhododen­ effervescing with enthusiasm about the mar­ dron. Numerous seepage areas and springs velous boreal bog that he had seen, and declar­ along the edge of the bog feed Sinking Creek. ing that he would make a special study of its Despite logging operations in the late 19th dragonflies and write a paper on it. For some century (3) and the annual invasion of blue­ reason he never did, and this present contribu­ berry pickers, Bear Meadows has remained tion is a monument to the spirit and character generally unaltered. In 1965 Bear Meadows of James G. Needham. was proclaimed a National Forest Monument PENNSYLVANIA ACADEMY OF SCIENCE 131

Figure 1. Bear Meadow;;, view from near the bridge looking west.

by Interior Secretary Udall, which should help RESULTS maintain this unusual habitat for future study The seasonal distribution, relative abundance and appreciation. and habitat affinities of the 58 species of PROCEDURE Odonata from Bear Meadows is summarized On most collecting trips to Bear Meadows it in Table 1. The fauna includes 7 species with was attempted to survey the species of Odon­ strong boreal affinities and 12 others with ata present and to estimate their relative weaker but definite boreal preferences. Ten abundance. A few sight records have been in­ species have a strong preference for streams cluded of species seen only on the wing but and are therefore associated with Sinking identified with certainty. Creek rather than the bog. Of the remaining Because of its ready access, the eastern end 29 species, 21 are frequent to common in cen­ of the bog adjacent to the bridge (Fig. 2) has tral Pennsylvania and do not show any spe­ received particular attention. This portion of cial preference for bog or stream environments. the bog contains a variety of sub-habitats and Eight species must be considered strays from is consequently populated by numerous species considerable distances. Their presence at Bear of Odonata. Surveys of the entire bog indicate Meadows is probably accidental. that collecting in the vicinity of the bridge is most productive and gives a reasonably repre­ Based on relative abundance, 29 or 50% of sentative sample of the species present. the species are rare. These are unlikely to be found during their flying season even when Data presented in this paper include those of N eedham's 1931 visit and several other early searched for. Twelve species are scarce, but collections, but over 80% of the records are the are usually collected in small numbers when result of work done since 1953. With the ex­ looked for. Ten species, observed consistently ception of data yielded by a collecting trip by without difficulty during their flying season, G. H. Beatty in June, 1945 (8) , none of the are termed common. The 7 species which are records in this paper has heretofore been re­ found in large numbers during their peak sea­ ported. son are classified as abundant. 132 PENNSYLVANIA ACADEMY OF SCIENCE

<:-<,.. BEAR MEADOWS . ~o CENTRE COUNTY, PENNSYLVANtf ELEVATION 1824 FEET ~

~BOG FOREST ~]BOG

1/4 MILE

·

0

DISCUSSION Somatochlora walshi Scudder Libellula quadrimaculata Linne Sphagnum bogs are among the less co=on Leucorrhinia hudsonica Selys habitat types in the Appalachians and their N ehalennia gracilis Morse Odonata faunas undoubtedly require much more study. This investigation of the Odonata In June, Gomphaeschna furcillata is an im­ of Bear Meadows has extended the known geo­ portant though not very conspicuous member graphical distribution of a number of species. of the Bear Meadows fauna. It is a cryptically It has also made the populations of other large colored species which blends with trunks of Pennsylvania bogs at Black Moshannon and dead trees on which it rests. The edge of the Tamarack, in Centre and Clinton Counties open meadow, especially where Sinking Creek respectively, more predictable since several of flows into the woods at the eastern end of the the species from Bear Meadows, which were bog, is the habitat of images of this species; unique records for the state, are now known the larvae have never been found at Bear from these other bogs. In the ensuing dis· Meadows though frequently sought. cussion, the Odonata population of Bear Mead­ Somatochlora elongata has been taken in few ows is divided into groups with similar ecologi­ Pennsylvania locations other than Bear Mead­ cal affinities. ows. This, however, is not the southernmost record since it has been reported from Mary­ The following species represent typical bog land (9), Virginia, and North Carolina (10). inhabitants and are part of the permanent Somatochlora elongata is not co=on at Bear breeding population. Meadows. Vlhen found, it is usually patrolling Gomphaeschna furcillata Say low over the stream, hovering frequently to ex­ Somatochlora elongata Scudder amine small coves along the bank, a flight Somatochlora tenebrosa Say pattern more characteristic of Aeshnids. PENNSYLVANIA ACADEMY OF SCIENCE 133

Since Bear Meadows is within the known probably part of the permanent breeding popu­ geographical range of Somatochlora tenebrosa, lation. its presence is of little distributional signifi­ Two species on this list are worthy of dis­ cance. It flies over the stream or hovers in cussion. Gomphus borealis has been collected sunlit clearings. on only 3 occasions. It is co=only and locally When Somatochlora walshi was first col­ abundant at Black Moshannon and Tamarack lected at Bear Meadows, it was the only Penn­ where it congregates on the bare ground, often sylvania occurrence and also the southern at the breast of beaver dams. Twice in this limit of its known distribution. Subsequently century beaver were introduced at Bear Mead­ it has been taken at the Black Moshannon and ows; however, they did not become established Tamarack bogs north of Bear Meadows but (3). Perhaps Gomphus borealis, along with still south of the Wisconsin Moraine. This spe­ the dead spires of drowned trees in the open cies is quite co=on some years. Occasionally meadow, is a vestige of short-lived beaver ac­ it flies over the stream, but more frequently tivity. it hovers over marshy pockets in the open Until recently (8) Epitheca canis was not meadow and sunny glades of the woodland known south of New York State. Studies in border. central Pennsylvania have shown this species The golden-flecked wings of Libellula quad­ to be relatively co=on and widespread al­ rimaculata are seen throughout June and July. though it has been taken only twice at Bear This typical bog species, circumpolar in distri­ Meadows. In central Pennsylvania this species bution, rests on the tips of twigs in the sun­ emerges before Epitheca cynosura. On May shine, flying out to capture small and 19, 1963 it was flying in great numbers in a returning to the same perch, a habit of many boggy area at the head of Stone Valley Dam Libellulas. Recreation Area in Huntingdon County. Males Leucorrhinia hudsonica is exceedingly abun­ patrolled territories 10 to 20 ft in diameter in dant at times. It alights on grass and sphag­ which they hovered, chasing away other males num moss near the stream and pools, or on of the same species. The frequency of this the ground of sunny woodland roads and clear­ species in Pennsylvania suggests that it will be ings. This species occurs at Black Moshannon discovered south of this state in the future. and Tamarack and also, irregularly at a few Another portion of the Odonata population other central Pennsylvania locations (11). at Bear Meadows is associated with the flowing Nehalennia gracilis is sometimes very abun­ water of Sinking Creek rather than the bog. dant, but could go unnoticed because of its These species are: small size and habit of flying and perching Basiaeschna janata Say low among the high marsh grasses. Boyeria vinosa Say In addition to the preceding species the fol­ Epiaeschna heros Fabricius lowing have boreal affinities but are less selec­ Aeshna umbrosa Walker tive in habitat requirements and therefore oc­ Cordulegaster disastatops Selys cur also in other kinds of habitats in central Cordulegaster maculatus Selys Pennsylvania. Lanthus parvulus Selys Aeshna canadensis Walker Helocordulia uhleri Selys Aeshna verticalis Hagen Calopteryx amata Hagen Gomphus borealis Needham Calopteryx maculata Beauvais Epitheca canis McLachlan Boyeria vinosa, Aeshna umbrosa, Cordule­ Cordulia shurtlefti Scudder gaster diastatops and Helocordulia uhleri are Libera Selys frequent to co=on most years and can be Libellula semifasciata Burmeister captured quite easily as they patrol up and Sympetrum obtrusum Hagen down the stream, Calopteryx maculata is often Sympetrum rubicundulum Say encountered along the stream where it rests Sympetrum semicinctum Say on overhanging grasses or sticks. It is most Chromagrion conditum Hagen co=on downstream from the bridge where Amphiagrion saucium Burmeister the current becomes swifter. Of these species only Cordulia shurtleffi, Basiaeschna janata, Cordulegaster macula­ Sympetrum rubicundulum and Chromagrion tus Lanthus parvulus and Calopteryx amata conditum are co=on every year. The others are probably strays from nearby streams where are sporadic in occurrence although they are they are frequently found. None of these has 134 PENNSYLVANIA ACADEMY OF SCIENCE been taken more than twice at Bear Meadows. Somatochlora linearis, Dorocordulia lepida, Epiaeschna heros has been seen on 4 occasions Libellula auripennis, and Libellula axilena are but never captured. It is probably an elusive not known from anywhere within about 100 but permanent resident of Bear Meadows. miles of Bear Meadows. These species have The remaining species are for the most part been collected only once. They are therefore, ubiquitous in central Pennsylvania, having not considered part of the permanent breeding either wide habitat tolerance or propensity for population. Archilestes grandis represents a wanderlust. noteworthy case. It was :first observed Septem­ ber 23, 1961. On the 19th of October that year Anax junius Drury it was quite co=on. The breeding population Epitheca cynosura Say appears to have been ephemeral since the only Libellula julia Uhler subsequent observation is a doubtful sight rec­ Libellula luctuosa Burmeister ord in 1962. The fact that this species flies late Libellula pulchella Drury in the year when collecting trips are infrequent Libellula lydia Drury may indicate that in most years Archilestes Erythemis simplicicollis Say grandis has been overlooked. Oddy enough, Pachydiplax longipennis Burmeister boreal Bear Meadows is one of the northern­ Leucorrhinia frigida Hagen most location for this species even though a Leucorrhinia intacta Hagen more austral environment might be expected as Sympetrum vicinum Hagen the northernmost station of a southern species. Lestes congener Hagen Lestes forcipatus Rambur In 1951 a collector visiting Bear Meadows Lestes rectangularis Say for the first time collected .specimens of Tacho­ Argia violace Hagen pteryx thoreyi, Dorocordulia lepida, Libellula N ehalennia irene Hagen cyanea and Enallagma ebrium. These species Enallagma civile Hagen have never been collected at Bear Meadows Enallagma ebrium Hagen before or since, although over 92 collecting trips Enallagma hageni Walsh have been made there. All but Dorocordulia Ischnura posita Hagen lepida are known from central Pennsylvania. Ischnura verticalis Say The specimens referred to have been examined With the exception of Sympetrum mcmum and are correctly identified. Certainly this is and Enallagma hageni which are abundant and 'an example of the unpredictability of collect­ Argia violacea which is co=on, the species ing results. on this list are scarce at. Bear Meadows al­ though all are co=on at other central Penn­ In the United States, at least, very few def­ sylvania locations. It seems likely, therefore, inite, circumscribable locations have had their that many of the individuals seen or collected Odonata faunas and populations studied con­ a·re strays from other habitats. This does not tinuously over a long period of time. Although exclude the possibility that they occasionally collections have admittedly been irregular, the complete their life cycle at Bear Meadows. In 30-odd years of study of Bear Meadows dragon­ some cases this is probable. flies yields a more comprehensive picture of what the fauna there actually consists of than Enallagma hageni is probably the most is available for any other location or environ­ abundant species at Bear Meadows possibly mental unit in North America. For as being exceeded by Nehalennia gracilis which vagile as Odonata it is only when comings is much less conspicuous. The former species and goings are averaged out over a consider­ occurs by the thousands during the peak of its able period of time, that the composition of the flying season. It is mainly found ski=ing over the water. truly resident fauna emerges, as indicated by more frequent collection-symbols on Table 1, The following do not fall into any of the while the sporadic or accidental occurrences preceeding categories: stand out as being of a quite different nature. Tachopteryx thoreyi Hagen Succession in populations goes on, Somatochlora linearis Hagen just as it does in the more easily observed and Dorocordulia lepida Hagen documented forest and vegetation composition, Libellula auripennis Burmeister and it is hoped that accumulation of future Libellula axilena Westwood data will make it possible to throw more light Archilestes grandis Rambur on successional stages in the Odonata fauna of Lestes dryas Kirby Bear Meadows. TABLE 1. The seasonal distribution, relative abundance, and habitat affinities of the odonata of Bear lv!eadows, Pennsylvania.

aJ Month lVIAY JUNE JULY AUGUST SEPTEMBER OCTOBER "''0 aJ R Quarterl 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 :5 ,g 1 ~ § Habitat affinity No. of collections 1 2 6 6 14 7 3 10 7 11 3 0 6 5 3 0 4 3 1 2 2 aJ..o ~ «$ 1. Gomphaeschna furcillata sx X X X X X X 3 sphagnum bogs 2. Basiaeschna janata X 1 streams 3. Boyeria vinosa X X X X-X X X-X 3 shady streams with rapids 4. Epiaeschna heros X-X X 1 small woodland streams 5. Aeshna canadensis X-X X X-X X X 2 boggy streams & lakes 6. --- umbrosa X X X X- X X X- X X X X X 3 woodland streams 7. --- verticalis X X X- X X X-X 2 boggy streams & lakes 8. Anax junius X X X X 1 still waters & slow streams i 9. Cordulegaster diastatops X X X X X X X 3 spring runs & brooks ~ 10. maculatus X 1 rapid woodland streams ~ 11. Tachopteryx thoreyi 4 X 1 spring bogs ~ 12. Lanthus parvulus X 1 small spring-fed streams ~ 13. Gomphus borealis X X X 1 beaver ponds, slow streams 14. Epitheca canis X 1 boggy ponds - streams 15. cynosura X -X X 1 ponds & streams ~ 16. Helocordulia uhleri X-X X 3 & 0 X -X rapid streams rivers qj 17. Somatochlora elongata X X X-XX 2 boggy streams & rivers r:n 18. linearis X 1 woodland streams 19. tenebrosa X X X-XX X--X 2 boggy streams g~ 20. walshi X X -X X X X-XX 3 I boggy streams 21. Cordulia shurtleffi X X X X X X X X 3 boggy ponds & streams 22. Dorocordulia lepida 4 X 1 I swamps & bogs 23. lib era X X -X X X 1 boggy ponds 24. Libellula auripennis X 1 I coastal waters, etc. 25. axilena X 1 coastal waters 26. cyanea4 X 1 standing water 27. julia X X X X 2 woodland ponds & marshes 28. luctuosa X 1 ponds 29. lydia X X X X X X X X X-X X 2 standing water 30. pulchella X X-X X-X X X-X X 2 ponds 31. quadrimaculata X X X X X X X X X X 3 sphagnum bogs 1-' 00 32. semifasciata X-X- X X-X 1 I marshy ponds <:n >--' TABLE 1. (continued) ... <:>:> m

""Q) Month MAY JUNE JULY AUGUST SEPTEMBER OCTOBER () ~ § Quarterl 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 ..... 'D Habitat affinity No. of collections 1 2 6 6 14 7 3 10 7 11 3 0 6 5 3 0 4 3 1 2 2 1 ] § p:;

40. hudsonica X X X X X X X X X 4 sphagnum bogs U1~ 41. intacta X- X X X X 2 ponds 42. Calopteryx amata X 1 swift woodland streams ~ 43. maculata X-X X X X X X X-XX X-X X 4 streams ~ 44. Archilestes grandis X- --X 1 small streams ;J> 45. Lestes congener X-XX --X 2 ponds 46. --- dryas 4 X 1 temporary pools, etc. 47. --- forcipatus X X 1 marshy ponds ~ ~ X X --X- X 1 woodland ponds 0 48. --- rectangularis >,j 49. Argia violacea X X X X X X X-XX 3 streams & lakes rn 50. Amphiagrion saucium X- X X -X 2 spring runs & bogs 51. N ehalennia gracilis X X- X X X X X X X-XX X-X 4 sphagnum bogs ~ 52. irene X X-X X X 2 swampy ponds g 53. Chromagrion conditum X X X X X X X X X 4 spring brooks & pools 54. Enallagma civile X X X 1 ponds, lakes, streams 55. ebrium4 X 1 marshy ponds & streams 56. hageni X X X X X X X X X-X X 4 boggy ponds, streams, etc. 57. Ischnura posita X X X 1 slow streams & ponds 58. verticalis X X X X -X X X X-X X X-X X 3 all standing waters No. of species observed 2 7 22 14 34 25 16 24 23 25 19 - 22 19 14- 13 6 2 3 5 0

1. Quarters refer to divisions of months: 1 - 1st-7th, 2 - 8th-15th, 3 - 16th-23rd, 4 - 24th-end. 2. Relative abundance: 1 - Rare, 2 - Scarce, 3 - Common, 4 - Abundant. Terms are defined in text. 3. An "X" indicates that species was collected or recorded during that quarter; A "-" indicates probable but unconfirmed presence of a species during that quarter. 4. Species reported by collectors other than the authors. PENNSYLVANIA ACADElVIY OF SCIENCE 137

ACKNOWLEDGMENTS 3. Westerfeld, W. F. 1959. Flora of Centre and Hunt· ingdon Counties with related historical, geological and This work is based in part on research par­ physiographic features. Penna. State Univ. Agri. E>.1J. tially supported by grants from. the Central Sta. Bull. 647. Fund for Research, Pennsylvania State Univer­ 4. ·westerfeld, W. F. 1961. Castanea 26: 1. sity, and the cost of publication has been paid 5. Kaniuka, R. P. 1964. New York Times 19 July, 1964. by the Department of Biology, Pennsylvania 6. Kovar, H. M. 1966. Frontiers 30:74. State University to which thanks are due. 7. Kovar, A. J. 1965. Penna. Acad. Sci. Proc. 38:16. 8. Beatty, G. H., III. 1946. Ent. News 57:1, 50, 76, 104. 9. Walker, E. M. 1925. The North American Dragonflies REFERENCES of the Genus Somatochlora. Univ. Toronto Stndies, 1. Leverett, P. 1934. Glacial deposits in Pennsylvania. Biol. Series 26. Penna. Geol. Surv. (4th series) Bull. G 7. 123 pp. 10. Byers, C. F. 1931. Ent. News 42:113. 2. Beatty, G. H. and A. F. 1968. Penna. Acad. Sci. 11. White, H. B., III. 1963. North Central Branch, Ent. Proc. 42:110. Soc. Amer. Proc. 18:120. ~.· ...... 11 I 6 .. PSC SP.y 0 UW .. 44 .4 tu. #Pi U EPAPSQ f4

r'I

THE ODONATA FAuNA OF BEAR MEADOWS ,

. A BOREAL BOG IN CENTRAt. PENNSYLVANIA

CORRECTIONS

p.l30, lst line of introduction: read 320 for 520

p.l30, 5th line of introduc;.tion: read "making it" for "the location of which makes it"

p.l36: insert, as footnote 5 to TABLE l, "Nomenclature and orde;r of taxa follow G.H. & A.F. Beatty's Check List and Bibliography of Pennsylvania Odonata (12)"

p.l37: add, at end of references, "12. Be~;!,:ty, G.H. & A.F. 1968. Penna. Acad. Sci. Proc. 42.:120."

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